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derived from knocked-out stem cells, and will therefore produce eggs or sperm containing the knocked-out gene. When these chimera mice are crossbred with others of the wild type, some of their offspring will have one copy of the knocked-out gene in all their cells. These mice do not retain any grey
253:, a gene that normal mice don't have and that confers resistance to a certain toxic agent (e.g., neomycin) or that produces an observable change (e.g. colour or fluorescence). In addition, a second gene, such as herpes tk+, is also included in the construct in order to accomplish a complete selection.
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Knocking out the activity of a gene provides information about what that gene normally does. Humans share many genes with mice. Consequently, observing the characteristics of knockout mice gives researchers information that can be used to better understand how a similar gene may cause or contribute
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in order to select for the transformations that occurred via homologous recombination. Any insertion of DNA that occurred via random insertion will die because they test positive for both the neomycin resistance gene and the herpes tk+ gene, whose gene product reacts with
Ganciclovir to produce a
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Another limitation is that conventional (i.e. non-conditional) knockout mice develop in the absence of the gene being investigated. At times, loss of activity during development may mask the role of the gene in the adult state, especially if the gene is involved in numerous processes spanning
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While knockout mouse technology represents a valuable research tool, some important limitations exist. About 15 percent of gene knockouts are developmentally lethal, which means that the genetically altered embryos cannot grow into adult mice. This problem is often overcome through the use of
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of a foster mother. This produces offspring that are either wildtype and coloured the same colour as the blastocyst donor (grey) or chimera (mixed) and partially knocked out. The chimera mice are crossed with a normal wildtype mouse (grey). This produces offspring that are either white and
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There is variability in the whole procedure depending largely on the strain from which the stem cells have been derived. Generally cells derived from strain 129 are used. This specific strain is not suitable for many experiments (e.g., behavioural), so it is very common to
321:: some parts of their bodies result from the original stem cells, other parts from the knocked-out stem cells. Their fur will show patches of white and grey, with white patches derived from the knocked-out stem cells and grey patches from the recipient blastocyst.
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Zan Y, Haag JD, Chen KS, Shepel LA, Wigington D, Wang YR, Hu R, Lopez-Guajardo CC, Brose HL, Porter KI, Leonard RA, Hitt AA, Schommer SL, Elegbede AF, Gould MN (June 2003). "Production of knockout rats using ENU mutagenesis and a yeast-based screening assay".
313:. For this example, we use blastocysts from a grey mouse. The blastocysts now contain two types of stem cells: the original ones (from the grey mouse), and the knocked-out cells (from the white mouse). These blastocysts are then implanted into the
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The embryonic stem cells that incorporated the knocked-out gene are isolated from the unaltered cells using the marker gene from step 1. For example, the unaltered cells can be killed using a toxic agent to which the altered cells are
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gene are associated with more than half of human cancers and often lead to tumours in a particular set of tissues. However, when the p53 gene is knocked out in mice, the animals develop tumours in a different array of tissues.
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in place of the original gene. The chances of a successful recombination event are relatively low, so the majority of altered cells will have the new sequence in only one of the two relevant chromosomes – they are said to be
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Knocking out a gene also may fail to produce an observable change in a mouse or may even produce different characteristics from those observed in humans in which the same gene is inactivated. For example, mutations in the
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but whose functions have not been determined. By causing a specific gene to be inactive in the mouse, and observing any differences from normal behaviour or physiology, researchers can infer its probable function.
398:. The confounding presence of neighbouring 129 genes on the knockout segment of genetic material has been dubbed the "flanking-gene effect". Methods and guidelines to deal with this problem have been proposed.
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is engineered which is very similar to the original gene and its immediate neighbour sequence, except that it is changed sufficiently to make the gene inoperable. Usually, the new sequence is also given a
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When these heterozygous offspring are interbred, some of their offspring will inherit the knocked-out gene from both parents; they carry no functional copy of the original unaltered gene (i.e. they are
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Another serious limitation is a lack of evolutive adaptations in knockout model that might occur in wild type animals after they naturally mutate. For instance, erythrocyte-specific coexpression of
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development. Conditional/inducible mutation approaches are then required that first allow the mouse to develop and mature normally prior to ablation of the gene of interest.
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heterozygous for the knocked out gene or grey and wildtype. White heterozygous mice can subsequently be crossed to produce mice that are homozygous for the knocked out gene.
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deadly toxin. Moreover, cells that do not integrate any of the genetic material test negative for both genes and therefore die as a result of poisoning with neomycin.
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for which the knockout technique can easily be applied. They are widely used in knockout experiments, especially those investigating genetic questions that relate to
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the offspring to other strains. Some genomic loci have been proven very difficult to knock out. Reasons might be the presence of repetitive sequences, extensive
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There are several thousand different strains of knockout mice. Many mouse models are named after the gene that has been inactivated. For example, the
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293:. Cells that were transformed with a vector containing the neomycin resistance gene and the herpes tk+ gene are grown in a solution containing
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Breeding scheme for producing knockout mice. Blastocysts containing cells, that are both wildtype and knockout cells, are injected into the
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knockout mouse is named after the p53 gene which codes for a protein that normally suppresses the growth of tumours by arresting
172:. Knockout mice also offer a biological and scientific context in which drugs and other therapies can be developed and tested.
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and blood cancers at an early age. Other mouse models are named according to their physical characteristics or behaviours.
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A laboratory mouse in which a gene affecting hair growth has been knocked out (left) is shown next to a normal lab mouse.
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Gerlai R (May 1996). "Gene-targeting studies of mammalian behavior: is it the mutation or the background genotype?".
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Montel-Hagen A, Kinet S, Manel N, Mongellaz C, Prohaska R, Battini JL, Delaunay J, Sitbon M, Taylor N (March 2008).
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671:, Lipp HP (July 2002). "Knockout mice: simple solutions to the problems of genetic background and flanking genes".
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There are several variations to the procedure of producing knockout mice; the following is a typical example.
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Examples of research in which knockout mice have been useful include studying and modeling different kinds of
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some of the electroporated stem cells will incorporate the new sequence with the knocked-out gene into their
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845:– A website for ordering embryonic stem cells, targeting vectors and transgenic mice generated by KMR.
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370:. In some instances, the gene may serve a different function in adults than in developing embryos.
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765:"Erythrocyte Glut1 triggers dehydroascorbic acid uptake in mammals unable to synthesize vitamin C"
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A detailed explanation of how knockout (KO) mice are created is located at the website of the
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112:. Aspects of the technology for generating knockout mice, and the mice themselves have been
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The
National Institutes of Health discusses some important limitations of this technique.
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and/or inducing apoptosis. Humans born with mutations that deactivate the p53 gene have
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A knockout mouse (left) that is a model for obesity, compared with a normal mouse
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The new sequence from step 1 is introduced into the stem cells from step 2 by
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Creating
Knockout Mice for Targeting Vector from Knockout Mice Research(KMR)
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839:– The website for ordering ES cells and mice generated by TIGM
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Millions of knockout mice are used in experiments each year.
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by replacing it or disrupting it with an artificial piece of
869:– community model organism database for the laboratory mouse
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The Knock Out Mouse
Project (KOMP) Data Coordination website
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Crusio WE, Goldowitz D, Holmes A, Wolfer D (February 2009).
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716:"Standards for the publication of mouse mutant studies"
606:. National Human Genome Research Institute. August 2015
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is much harder and has only been possible since 2003.
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The Knock Out Mouse
Project (KOMP) Repository website
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List of varieties of genetically modified maize/corn
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The gene to be knocked out is isolated from a mouse
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Texas A&M Institute for
Genomic Medicine (TIGM)
219:The procedure for making mixed-genotype blastocyst
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93:The first recorded knockout mouse was created by
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583:"The Nobel Prize in Physiology or Medicine 2007"
27:Genetically modified mouse used in gene research
62:for studying the role of genes which have been
851:– a review from the Science Creative Quarterly
849:Studying Gene Function: Creating Knockout Mice
105:in 1989, for which they were awarded the 2007
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46:) in which researchers have inactivated, or "
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362:. The lack of adult mice limits studies to
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324:Some of the newborn chimera mice will have
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564:. National Human Genome Research Institute
452:International Mouse Phenotyping Consortium
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585:. Nobelprize.org. 1985-09-19
280:. By the natural process of
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560:Spencer G (December 2002).
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604:"Knockout Mice Fact Sheet"
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720:Genes, Brain and Behavior
480:Pilcher HR (2003-05-19).
110:in Physiology or Medicine
72:laboratory animal species
1720:Laboratory mouse strains
282:homologous recombination
74:most closely related to
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626:Trends in Neurosciences
70:Mice are currently the
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1040:
1038:
1034:
1028:
1025:
1023:
1020:
1019:
1017:
1015:
1011:
1005:
1002:
1001:
999:
997:
993:
987:
984:
983:
981:
979:
975:
969:
966:
964:
961:
959:
956:
954:
951:
950:
948:
946:
942:
939:
937:
933:
930:
928:
920:
916:
909:
904:
902:
897:
895:
890:
889:
886:
879:
876:
874:
871:
868:
865:
862:
859:
856:
853:
850:
847:
844:
841:
838:
835:
834:
827:
824:
822:
819:
818:
813:
808:
792:
788:
783:
778:
774:
770:
766:
759:
751:
747:
743:
739:
734:
729:
725:
721:
717:
710:
702:
698:
694:
690:
686:
682:
679:(7): 336–40.
678:
674:
670:
663:
655:
651:
647:
643:
639:
635:
632:(5): 177–81.
631:
627:
620:
605:
599:
584:
578:
563:
556:
554:
545:
541:
537:
533:
529:
525:
522:(6): 645–51.
521:
517:
509:
495:
491:
487:
483:
476:
472:
463:
460:
458:
457:Knockout moss
455:
453:
450:
448:
445:
443:
440:
438:
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430:
428:
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296:
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283:
279:
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272:
271:
266:
262:
258:
255:
252:
247:
244:. Then a new
243:
239:
238:
237:
230:
225:
217:
208:
206:
205:breast cancer
202:
198:
197:cell division
194:
185:
176:
173:
171:
167:
163:
159:
155:
151:
147:
146:heart disease
143:
139:
134:
126:
117:
115:
111:
108:
104:
100:
96:
91:
89:
85:
84:Gene knockout
81:
77:
73:
68:
65:
61:
60:animal models
57:
53:
49:
45:
41:
37:
33:
19:
18:Knockout mice
1478:Gene therapy
1377:Transgenesis
1356:Transfection
1321:Agrobacteria
1205:Knockout rat
1182:
1027:Vistive Gold
923:Genetically
811:
772:
768:
758:
723:
719:
709:
676:
672:
662:
629:
625:
619:
608:. Retrieved
598:
587:. Retrieved
577:
566:. Retrieved
519:
515:
508:
497:. Retrieved
485:
475:
404:
400:
384:
374:
368:human health
356:
352:
340:
291:heterozygous
268:
267:) and grown
246:DNA sequence
242:gene library
235:
190:
174:
135:
131:
99:Martin Evans
92:
69:
44:Mus musculus
43:
35:
31:
29:
1497:In research
1471:diagnostics
1341:Lipofection
1102:Arabidopsis
1004:Golden rice
667:Wolfer DP,
349:Limitations
299:Ganciclovir
286:chromosomes
251:marker gene
107:Nobel Prize
48:knocked out
1709:Categories
1581:Regulation
1425:Companies
1382:Cisgenesis
1326:Biolistics
1132:Sugar beet
1045:Flavr Savr
945:Maize/corn
726:(1): 1–4.
610:2014-04-03
589:2014-04-03
568:2014-04-03
499:2014-04-03
468:References
335:homozygous
311:blastocyst
306:resistant.
261:blastocyst
1694:Bioethics
1598:Geography
1536:Transgene
1304:Processes
1195:Enviropig
1188:Oncomouse
1127:SmartStax
1063:Bt cotton
927:organisms
750:205853147
669:Crusio WE
462:Oncomouse
415:vitamin C
388:backcross
211:Procedure
154:arthritis
64:sequenced
1684:Genetics
1651:Eugenics
1529:articles
1454:Syngenta
1449:Monsanto
1421:Pharming
1267:Bacteria
963:StarLink
925:modified
791:18358815
742:18778401
701:33777888
693:12079755
654:33396039
544:32611710
536:12754522
437:Genetics
421:See also
411:stomatin
319:chimeras
295:neomycin
270:in vitro
150:diabetes
114:patented
1679:Biology
1656:Cloning
1635:Oceania
1527:Related
1272:viruses
1236:GloFish
1226:Insects
1170:Mammals
1161:Animals
1147:Mustard
1137:Tobacco
1107:Brinjal
1014:Soybean
986:Amflora
958:MON 863
953:MON 810
646:8723200
442:Humouse
179:Strains
162:anxiety
142:obesity
38:, is a
1610:Africa
1605:Europe
1467:humans
1241:Salmon
1210:Rabbit
1117:Papaya
1112:Canola
1055:Cotton
1037:Tomato
978:Potato
809:about
789:
748:
740:
699:
691:
652:
644:
542:
534:
486:Nature
345:2007.
326:gonads
315:uterus
265:embryo
229:uterus
138:cancer
101:, and
76:humans
1434:Bayer
1365:Types
1253:Frogs
1248:Birds
1178:Mouse
1142:Trees
1097:Apple
1090:Other
1073:Wheat
936:Crops
746:S2CID
697:S2CID
650:S2CID
540:S2CID
409:with
407:GLUT1
394:, or
166:aging
34:, or
1615:Asia
1429:BASF
1395:Uses
1270:and
1231:Fish
1122:Rose
996:Rice
787:PMID
769:Cell
738:PMID
689:PMID
642:PMID
532:PMID
297:and
168:and
88:rats
52:gene
1469:and
1465:In
1081:HB4
777:doi
773:132
728:doi
681:doi
634:doi
524:doi
490:doi
378:p53
193:p53
120:Use
86:in
56:DNA
1711::
1624:US
785:.
771:.
767:.
744:.
736:.
722:.
718:.
695:.
687:.
677:25
675:.
648:.
640:.
630:19
628:.
552:^
538:.
530:.
520:21
518:.
488:.
484:.
417:.
164:,
160:,
156:,
152:,
148:,
144:,
140:,
97:,
82:.
30:A
1626:)
1622:(
907:e
900:t
893:v
793:.
779::
752:.
730::
724:8
703:.
683::
656:.
636::
613:.
592:.
571:.
546:.
526::
502:.
492::
42:(
20:)
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